268:. The action of these insulators is dependent on being positioned between the promoter of the target gene and the upstream or down stream enhancer. The specific way in which insulators block enhancers is dependent on the enhancers mode of action. Enhancers can directly interact with their target promoters through looping (direct-contact model), in which case an insulator prevents this interaction through the formation of a loop domain that separates the enhancer and promoter sites and prevents the promoter-enhancer loop from forming. An enhancer can also act on a promoter through a signal (tracking model of enhancer action). This signal may be blocked by an insulator through the targeting of a nucleoprotein complex at the base of the loop formation.
253:
1445:
427:
CTCF was first characterized for its role in regulating β-globin gene expression. At this locus, CTCF functions as an insulator-binding protein forming a chromosomal boundary. CTCF is present in both the chicken β-globin locus and human β-globin locus. Within cHS4 of the chicken β-globin locus, CTCF
453:
imprinted locus. In this locus the CTCF protein regulates imprinted expression by binding to the unmethylated maternal imprinted control region (ICR) but not on the paternal ICR. When bound to the unmethylated maternal sequence, CTCF effectively blocks downstream enhancer elements from interacting
259:
Similar mechanism of action for enhancer-blocking insulators; chromatin loop domains are formed in the nucleus that separates the enhancer and the promoter of a target gene. Loop domains are formed through the interaction between enhancer-blocking elements interacting with each other or securing
276:
Barrier activity has been linked to the disruption of specific processes in the heterochromatin formation pathway. These types of insulators modify the nucleosomal substrate in the reaction cycle that is central to heterochromatin formation. Modifications are achieved through various mechanisms
330:
imprinted locus where methylation of the paternal imprinted control region (ICR) prevents CTCF from binding. A second mechanism of regulation is through regulating proteins that are required for fully functioning CTCF insulators. These proteins include, but are not limited to
297:
insulator appears to have enhancer blocking activity via its 3D structure and have no direct connection with barrier activity. Vertebrates in particular appear to rely heavily on the CTCF insulator, however there are many different insulator sequences identified.
552:, which are growth suppressors that are silenced in many cancer types, and whose expression is controlled by CTCF. Loss of function of CTCF in these genes leads to the silencing of the growth suppressor and contributes to the formation of cancer.
482:
of a gene, it has been suggested that they might serve to stabilize enhancer-promoter interactions. When they are located farther away from the promoter, insulator elements would compete with the enhancer and interfere with activation of
384:
that are both tissue specific and present at certain developmental stages. The insulator likely has an inhibitory effect on enhancers that control the spatial and temporal expression of the affected gene.
491:) into closer proximity for interaction during transcription. The mechanism of enhancer-blocking insulators then, if in the correct position, could play a role in regulating transcription activation.
163:
While enhancer-blocking is classified as an inter-chromosomal interaction, acting as a barrier is classified as an intra-chromosomal interaction. The need for insulators arises where two adjacent
420:. Different from the chicken β-globin locus, the human β-globin locus has an open chromatin structure and is not flanked by a 5' heterochromatic region. HS5 is thought to be a genetic insulator
405:
insulator acts as both a barrier to chromatin-mediated silencing via heterochromatin spreading, and blocks interactions between enhancers and promoters. A distinguishing characteristic of
281:
removal, in which nucleosome-excluding elements disrupt heterochromatin from spreading and silencing (chromatin-mediated silencing). Modification can also be done through recruitment of
175:
patterns; it is critical that the inducing or repressing mechanisms of one do not interfere with the neighbouring gene. Insulators have also been found to cluster at the boundaries of
523:
mutation to the CTCF insulator gene changes the expression patterns and may affect the interplay between cell growth, differentiation and apoptosis and lead to
821:
Wai, Albert W.K.; Gillemans, Nynke; Raguz-Bolognesi, Selina; Pruzina, Sara; Zafarana, Gaetano; Meijer, Dies; Philipsen, Sjaak; Grosveld, Frank (2003-09-01).
401:
marks the border between the active euchromatin in the β-globin locus and the upstream heterochromatin region that is highly condensed and inactive. The
182:
Some insulators can act as both enhancer blocker and barriers, and some just have one of the two functions. Some examples of different insulators are:
1477:
1623:
1186:
Dowen, JM; Fan, ZP; Hnisz, D; Ren, G; Abraham, BJ; Zhang, LN; Weintraub, AS; Schuijers, J; Lee, TI; Zhao, K; Young, RA (9 October 2014).
179:(TADs) and may have a role in partitioning the genome into "chromosome neighborhoods" - genomic regions within which regulation occurs.
1391:"Chromatin insulation orchestrates matrix metalloproteinase gene cluster expression reprogramming in aggressive breast cancer tumors"
1249:
642:
588:
302:
formed by physical interaction between two CTCF-bound DNA loci contain the interactions between enhancers and their target genes.
176:
1827:
878:
Burgess-Beusse B, Farrell C, Gaszner M, Litt M, Mutskov V, Recillas-Targa F, Simpson M, West A, Felsenfeld G (December 2002).
1449:
79:-blocker or a barrier, or both. The mechanisms by which an insulator performs these two functions include loop formation and
110:
The β-globin locus was first studied in chicken and then in humans for its insulator activity, both of which utilize CTCF.
1470:
27:
1503:
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1606:
1584:
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1920:
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219:
172:
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1490:
484:
187:
118:
20:
1710:
1463:
823:"HS5 of the human β-globin locus control region: a developmental stage-specific border in erythroid cells"
299:
46:
38:
1033:"Controlling long-range genomic interactions at a native locus by targeted tethering of a looping factor"
1894:
1889:
1757:
1659:
475:
323:
541:. When the CTCF binding site is removed expression of Rb is decreased and tumours are able to thrive.
1705:
1671:
1565:
1335:
891:
555:
The aberrant activation of insulators can modulate the expression of cancer-related genes, including
488:
65:
318:. CTCF protein is known to favourably bind to unmethylated sites, so it follows that methylation of
1774:
1742:
1628:
1602:
479:
365:
146:
76:
53:
1543:
1353:
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114:
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Deng, W; Lee, J; Wang, H; Miller, J; Reik, A; Gregory, P. D.; Dean, A; Blobel, G. A. (2012).
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772:
708:
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520:
1188:"Control of cell identity genes occurs in insulated neighborhoods in mammalian chromosomes"
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463:
369:
157:
487:. Loop formation is common in eukaryotes to bring distal elements (enhancers, promoters,
1455:
1417:
1390:
1389:
Llinàs-Arias P, Ensenyat-Mendez M, Íñiguez-Muñoz S, Orozco J, Valdez B (November 2023).
1339:
895:
393:
The first examples of insulators in vertebrates was seen in the chicken β-globin locus,
1858:
1846:
1796:
1299:
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1212:
1187:
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336:
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1914:
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1366:
1323:
979:
511:). Two of these cell cycle regulation genes that are known to interact with CTCF are
203:
61:
800:
16:
Genetic boundary element that blocks the interaction between enhancers and promoters
315:
265:
231:
113:
The genetic implications of insulators lie in their involvement in a mechanism of
1747:
1715:
1618:
446:
319:
311:
153:
92:
1407:
1328:
Proceedings of the
National Academy of Sciences of the United States of America
1324:"Genetic and Molecular Analysis of the gypsy Chromatin Insulator of Drosophila"
1203:
1146:
1097:
1048:
1899:
1593:
1548:
1538:
1533:
1528:
1523:
538:
500:
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168:
126:
105:
96:
80:
69:
1290:
1154:
846:
784:
722:
652:
1555:
1348:
1267:"CTCF as a multifunctional protein in genome regulation and gene expression"
508:
428:
binds to a region (FII) that is responsible for enhancer blocking activity.
424:
as it has both enhancer-blocking activity and transgene barrier activities.
381:
261:
57:
49:
1426:
1308:
1221:
1172:
1115:
1066:
1017:
964:
923:
904:
864:
838:
792:
730:
670:
252:
1444:
1375:
619:"Chromatin Insulators: A Role in Nuclear Organization and Gene Expression"
1762:
1752:
880:"The insulation of genes from external enhancers and silencing chromatin"
535:
1282:
1722:
1589:
713:
696:
534:(Rb) gene and mutations and deletions of this gene are associated with
332:
984:"CTCF: an architectural protein bridging genome topology and function"
955:
938:
376:
affects the expression of adjacent genes pending insertion into a new
1518:
1357:
377:
122:
999:
776:
760:
83:
modifications. There are many examples of insulators, including the
1322:
Gdula, David A.; Gerasimova, Tatiana I.; Corces, Victor G. (1996).
1779:
1631:
761:"Insulators: exploiting transcriptional and epigenetic mechanisms"
545:
409:
is that it has a repetitive heterochromatic region on its 5' end.
360:
is one of several sequences that have been studied in detail. The
211:
that have both enhancer-blocking and barrier activity, as well as
1850:
1680:
1611:
294:
164:
84:
1459:
695:
West, Adam G.; Gaszner, Miklos; Felsenfeld, Gary (2002-02-01).
549:
530:
CTCF is also required for the expression of tumour repressor
499:
CTCF insulators affect the expression of genes implicated in
583:. New Jersey: John Wiley & Sons, Inc. pp. 300–301.
251:
445:
genes is dependent on the presence of an insulator on the
56:
element of the target gene, an insulator is typically 300
503:
regulation processes that are important for cell growth,
1265:
Kim, Somi; Yu, Nam-Kyung; Kaang, Bong-Kiun (June 2015).
449:
allele between the two genes. An example of this is the
285:(s) and ATP-dependent nucleosome remodelling complexes.
1244:. New Jersey: John Wiley & Sons, Inc. p. 367.
1129:
Phillips, Jennifer E.; Corces, Victor G. (2009-06-26).
544:
Other genes that encode cell cycle regulators include
95:
locus. The CTCF insulator is especially important in
759:
Gaszner, Miklos; Felsenfeld, Gary (September 2006).
1871:
1836:
1810:
1735:
1693:
1670:
1647:
1583:
1576:
1511:
1502:
60:to 2000 bp in length. Insulators contain clustered
149:from acting on the promoter of neighbouring genes
1471:
697:"Insulators: many functions, many mechanisms"
8:
145:Enhancer-blocking insulators prevent distal
350:The insulator element that is found in the
1580:
1508:
1478:
1464:
1456:
617:Yang, Jingping; Corces, Victor G. (2011).
1416:
1406:
1365:
1347:
1298:
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1162:
1105:
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1007:
954:
913:
903:
854:
712:
660:
215:that have only enhancer-blocking activity
156:by inhibiting the spread of neighbouring
310:One mechanism of regulating CTCF is via
264:fibre to structural elements within the
152:Barrier insulators prevent silencing of
121:. Mutations to insulators are linked to
568:
133:, and silencing of growth suppressors.
478:are located in close proximity to the
412:The human β-globin locus homologue of
1624:Histone acetylation and deacetylation
1271:Experimental & Molecular Medicine
1235:
1233:
1231:
441:The ability of enhancers to activate
238:insulator acts as an enhancer-blocker
199:are both enhancer-blocking insulators
7:
1080:Phillips JE, Corces VG (June 2009).
816:
814:
812:
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754:
752:
750:
748:
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684:
682:
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612:
610:
608:
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602:
600:
574:
572:
326:. An example of this is seen in the
141:Insulators have two main functions:
52:and working over distances from the
1131:"CTCF: Master Weaver of the Genome"
1082:"CTCF: master weaver of the genome"
939:"Mapping chromosome neighborhoods"
635:10.1016/B978-0-12-386469-7.00003-7
559:involved in cancer cell invasion.
14:
364:insulator can be found in the 5'
177:topologically associating domains
75:Insulators function either as an
1443:
458:gene promoter, leaving only the
1828:Archaeal transcription factor B
117:and their ability to regulate
19:For electrical insulator, see
1:
1242:Fundamental Molecular Biology
1240:Allison, Lizabeth A. (2012).
581:Fundamental Molecular Biology
579:Allison, Lizabeth A. (2012).
507:, and programmed cell death (
68:and mediate intra- and inter-
884:Proc. Natl. Acad. Sci. U.S.A
248:Enhancer-blocking insulators
623:Advances in Cancer Research
227:are both barrier insulators
103:insulator is implicated in
28:Insulation (disambiguation)
1939:
1504:Transcriptional regulation
1408:10.1186/s12943-023-01906-8
1204:10.1016/j.cell.2014.09.030
1147:10.1016/j.cell.2009.06.001
1098:10.1016/j.cell.2009.06.001
1049:10.1016/j.cell.2012.03.051
25:
18:
1701:Transcription coregulator
1637:Histone acetyltransferase
1607:Histone methyltransferase
1585:Histone-modifying enzymes
557:matrix metalloproteinases
380:location, causing mutant
283:histone acetyltransferase
937:Perkel J (1 June 2015).
220:Saccharomyces cerevisiae
1802:Internal control region
1349:10.1073/pnas.93.18.9378
765:Nature Reviews Genetics
701:Genes & Development
300:Insulated neighborhoods
223:insulators STAR and UAS
188:Drosophila melanogaster
21:Insulator (electricity)
905:10.1073/pnas.162342499
256:
64:for sequence specific
41:known as a long-range
39:cis-regulatory element
1895:Intrinsic termination
1660:DNA methyltransferase
489:locus control regions
324:epigenetic regulation
255:
1672:Chromatin remodeling
1452:at Wikimedia Commons
1450:Insulator (genetics)
890:(Suppl 4): 16433–7.
839:10.1093/emboj/cdg437
505:cell differentiation
495:Mutations and cancer
432:Genetic implications
171:have very different
66:DNA-binding proteins
26:For other uses, see
1629:Histone deacetylase
1619:Histone demethylase
1603:Histone methylation
1340:1996PNAS...93.9378G
1283:10.1038/emm.2015.33
896:2002PNAS...9916433B
527:or other problems.
366:untranslated region
354:retrotransposon of
243:Mechanism of action
91:insulator, and the
714:10.1101/gad.954702
519:In these cases, a
272:Barrier insulators
257:
43:regulatory element
1908:
1907:
1863:RNA polymerase II
1731:
1730:
1689:
1688:
1448:Media related to
1334:(18): 9378–9383.
956:10.2144/000114296
833:(17): 4489–4500.
207:have insulators,
1928:
1785:Response element
1768:Response element
1581:
1509:
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1395:Molecular Cancer
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827:The EMBO Journal
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521:loss of function
1938:
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1649:DNA methylation
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1092:(7): 1194–211.
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1000:10.1038/nrg3663
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777:10.1038/nrg1925
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474:When insulator
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370:retrotransposon
348:
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158:heterochromatin
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129:disregulation,
125:as a result of
87:insulator, the
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1859:RNA polymerase
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1847:RNA polymerase
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1438:External links
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1043:(6): 1233–44.
1023:
982:(April 2014).
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771:(9): 703–713.
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707:(3): 271–288.
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532:retinoblastoma
525:tumourigenesis
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389:β-globin locus
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337:RNA polymerase
322:is a point of
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289:CTCF insulator
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1566:trp repressor
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1561:lac repressor
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1487:Transcription
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1198:(2): 374–87.
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62:binding sites
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47:multicellular
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539:malignancies
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447:unmethylated
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316:DNA sequence
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232:Homo sapiens
230:
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100:
99:, while the
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74:
34:
32:
1872:Termination
1748:Pribnow box
1716:Corepressor
1711:Coactivator
1512:prokaryotic
1277:(6): e166.
462:gene to be
320:CpG islands
312:methylation
191:insulators
154:euchromatin
97:vertebrates
70:chromosomal
45:. Found in
1900:Rho factor
1890:Terminator
1881:eukaryotic
1856:eukaryotic
1837:Elongation
1823:Eukaryotic
1811:Initiation
1594:nucleosome
1577:eukaryotic
1549:gal operon
1544:ara operon
1539:Gua Operon
1534:gab operon
1529:trp operon
1524:lac operon
1495:Eukaryotic
1401:(4): 190.
563:References
501:cell cycle
437:Imprinting
382:phenotypes
357:Drosophila
306:Regulation
279:nucleosome
277:including
169:chromosome
127:cell cycle
115:imprinting
106:Drosophila
81:nucleosome
50:eukaryotes
1876:bacterial
1844:bacterial
1818:Bacterial
1792:Insulator
1736:Promotion
1706:Activator
1556:Repressor
1491:Bacterial
1291:2092-6413
1155:0092-8674
980:Corces VG
847:0261-4189
785:1471-0064
723:0890-9369
653:0065-230X
629:: 43–76.
536:inherited
509:apoptosis
476:sequences
464:expressed
454:with the
443:imprinted
372:element.
346:insulator
262:chromatin
147:enhancers
35:insulator
1915:Category
1797:Silencer
1775:Enhancer
1763:CAAT box
1753:TATA box
1743:Promoter
1427:38017545
1418:10683115
1309:26045254
1222:25303531
1173:19563753
1116:19563753
1067:22682246
1018:24614316
978:Ong CT;
965:26054763
924:12154228
865:12941700
801:31291034
793:16909129
731:11825869
671:21704228
480:promoter
451:Igf2-H19
328:Igf2-H19
209:Lys 5' A
137:Function
93:β-globin
77:enhancer
54:promoter
1723:Inducer
1590:histone
1376:8790337
1336:Bibcode
1300:4491725
1213:4197132
1164:3040116
1107:3040116
1058:3372860
1009:4610363
892:Bibcode
662:3175007
422:in vivo
378:genomic
333:cohesin
314:of its
266:nucleus
197:scs scs
1519:Operon
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548:, and
517:C-MYC.
123:cancer
1780:E-box
1632:HDAC1
1367:38435
1358:39717
1354:JSTOR
797:S2CID
546:BRCA1
513:hTERT
374:Gypsy
362:gypsy
352:gypsy
344:gypsy
193:gypsy
167:on a
165:genes
101:gypsy
89:gypsy
1851:rpoB
1694:both
1681:CHD7
1612:EZH2
1423:PMID
1372:PMID
1305:PMID
1287:ISSN
1246:ISBN
1218:PMID
1192:Cell
1169:PMID
1151:ISSN
1135:Cell
1112:PMID
1086:Cell
1063:PMID
1037:Cell
1014:PMID
961:PMID
920:PMID
861:PMID
843:ISSN
789:PMID
781:ISSN
727:PMID
719:ISSN
667:PMID
649:ISSN
639:ISBN
585:ISBN
515:and
456:Igf2
414:cHS4
407:cHS4
403:cHS4
399:cHS4
395:cHS4
295:CTCF
293:The
195:and
85:CTCF
1758:BRE
1413:PMC
1403:doi
1362:PMC
1344:doi
1295:PMC
1279:doi
1208:PMC
1200:doi
1196:159
1159:PMC
1143:doi
1139:137
1102:PMC
1094:doi
1090:137
1053:PMC
1045:doi
1041:149
1004:PMC
996:doi
951:doi
910:PMC
900:doi
851:PMC
835:doi
773:doi
709:doi
657:PMC
631:doi
627:110
550:p53
460:H19
418:HS5
416:is
236:HS5
225:rpg
213:HS4
33:An
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